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Full text of "A key to the bats of the Philippine Islands"

IBUKY UBRARY 
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FIELDIANA 



Zoology 

NEW SERIES, NO. 69 



A Key to the Bats of the Philippine Islands 

Nina R. Ingle 

Department of Natural Resources 
Fernow Hall 
Cornell University 
Ithaca, New York 14853 

Lawrence R. Heaney 

Department of Zoology 
Field Museum of Natural History 
Roosevelt Road at Lake Shore Drive 
Chicago. Illinois 60605-2496 



Accepted March 20, 1992 
Published October 30, 1992 
Publication 1440 



PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY 









© 1992 Field Museum of Natural History 
Library of Congress Catalog Card Number: 92-73951 

,m ISSN 0015-0754 

PRINTED IN THE UNITED STATES OF AMERICA 



Table of Contents 



Abstract 1 

Introduction 1 

Methods 3 

Anatomy and Measurements of a Bat .... 3 

Use of the Key 4 

Determination of Age and Sex 5 

Identification to Family and Species 6 

Use of Measurement Tables 6 

Comparison with Skull Drawings 6 

Collecting and Preserving Voucher Spec- 
imens 7 

Key to Families of Philippine Bats 7 

Family Pteropodidae: Fruit Bats 8 

Key to Pteropodidae 10 

Family Emballonuridae: Sheath-Tailed 

Bats 14 

Key to Emballonuridae 14 

Family Megadermatidae: False Vampire 

Bats 14 

Family Rhinolophidae: Horseshoe and 

Leaf-Nosed Bats 15 

Key to Rhinolophidae 15 

Family Vespertilionidae: Evening Bats .18 

Key to Vespertilionidae 20 

Family Molossidae: Free-Tailed Bats .... 22 

Key to Molossidae 23 

Notes 23 

Acknowledgments 25 

Literature cited 25 

List of Illustrations 

1. Map of the Philippines 2 

2. Generalized pteropodid bat 3 

3. Ears of representative bats 4 

4. Bat with external measurements indicat- 
ed 4 

5. Bat skull with teeth and cranial mea- 
surements indicated 5 

6. Joints in digits of wing of adult and ju- 
venile bats 5 

7. Tails of representatives of Philippine 

bat families 8 

8. Heads of representatives of Philippine 

bat families 9 

9. Frontal views of fruit bat skulls, show- 
ing incisors and canines 12 

10. Dorsal views of fruit bat muzzles 13 

1 1 . Posterior attachment of wing membrane 

on fruit bats 14 



1 2. Noseleaves of Rhinolophus and Hipposi- 
deros 16 

1 3. Connecting processes of some rhinolo- 
phids 18 

14. Ventral views of muzzles of Murina and 
Myotis 21 

1 5. External ears (pinnae) of some vespertil- 
ionids 21 

16. Hind feet of Myotis spp., showing at- 
tachment of wing membrane 22 

1 7. Pad on wrist of Glischropus 22 

1 8. Ears of Tylonycteris 22 

19. Skull of Acerodon jubatus 28 

20. Skull of Cynopterus brachyotis 29 

2 1 . Skull of Dobsonia chapmani 29 

22. Skull of Eonycteris robusta 29 

23. Skull of Eonycteris spelaea 30 

24. Skull of Haplonycteris fischeri 30 

25. Skull of Harpyionycteris whiteheadi .... 30 

26. Skull of Macroglossus minimus 31 

27. Skull of Nyctimene rabori 31 

28. Skull of Ptenochirus jagori 31 

29. Skull of Pteropus hypomelanus 32 

30. Skull of Pteropus pumilus 32 

3 1 . Skull of Pteropus vampyrus 33 

32. Skull of Rousettus ample xicaudatus .... 33 

33. Skull of Emballonura alecto 33 

34. Skull of Saccolaimus saccolaimus 34 

35. Skull of Taphozous melanopogon 34 

36. Skull of Megaderma spasma 35 

37. Skull of Hipposideros ater 35 

38. Skull of Hipposideros diadema 36 

39. Skull of Hipposideros obscurus 36 

40. Skull of Hipposideros pygmaeus 37 

41. Skull of Rhinolophus arcuatus-X 37 

42. Skull of Rhinolophus arcuatus-s 38 

43. Skull of Rhinolophus inops 38 

44. Skull of Rhinolophus macrotis 39 

45. Skull of Rhinolophus philippinensis .... 39 

46. Skull of Rhinolophus virgo 40 

47. Skull of Kerivoula hardwickii 40 

48. Skull of Miniopterus australis 40 

49. Skull of Miniopterus schreibersi 41 

50. Skull of Miniopterus tristis 41 

5 1 . Skull of Murina cyclotis 41 

52. Skull of Myotis horsfieldii 42 

53. Skull of Myotis macrotarsus 42 

54. Skull of Myotis muricola 42 

55. Skull of Myotis rufopictus 43 

56. Skull of Philetor brachypterus 43 

57. Skull of Pipistrellus javanicus 43 

58. Skull of Pipistrellus tenuis 44 

59. Skull of Scotophilus kuhlii 44 

60. Skull of Chaerephon plicata 44 



in 



List of Tables 



1. Measurements of Philippine Pteropodi- 

dae 11 

2. Measurements of Philippine Emballonur- 
idae and Megadermatidae 15 

3. Measurements of Philippine Rhinolophi- 
dae 17 

4. Measurements of Philippine Vespertilion- 
idae 19 

5. Measurements of Philippine Molossidae. . 24 






IV 



A Key to the Bats of the Philippine Islands 

Nina R. Ingle and Lawrence R. Heaney 



Abstract 

An identification guide is presented for the six families and 70 species of bats now known 
from the Philippine Islands, based on a key and a set of standardized measurements. Most 
critical characters are illustrated, and detailed drawings are provided of the skulls of 42 species. 



Introduction 

The Philippine Islands (fig. 1) support a large 
and diverse fauna of mammals: over 1 70 species 
are now known, compared, for example, to the 
105 species known from Madagascar, which has 
nearly twice the land area (Heaney et al., 1987; 
Jenkins, 1987). About 100 species are endemic to 
the Philippines, giving the country an unusually 
high number and percentage of unique species 
(HaugeetaL, 1986; Heaney, 1986, 1991; Heaney 
et al., 1987; Koopman, 1989). 

One of the most diverse and, in general, poorly 
known mammalian orders in the Philippines is 
the Chiroptera. Sixty-eight species of bats were 
known from the Philippines when the last checklist 
was prepared (Heaney et al., 1987), and two more 
are now known (Pteropus dasymallus and Harpi- 
ocephalus harpia). In number of species, bats ex- 
ceed even rodents, of which 67 are now known 
(Heaney et al., 1987; Musser and Heaney, 1992). 
By our best estimate, 22 species of bats, about 
31%, are endemic to the Philippines, again an un- 
usually high number (Heaney, 1991; Koopman, 
1989). 

The high levels of species richness and ende- 
mism are factors of special importance currently 
because of the rapid rate of loss of natural habitat 
in the Philippines. Roughly 94% of the Philippine 
land area was once covered by forest; that figure 
had been reduced to 40% at the end of World War 
II, and current estimates of forest cover range from 
25% to less than 20%, depending in part on the 
amount of degraded forest that is included (Col- 
lins, 1990; Hauge et al., 1986; Myers, 1988; Ut- 
zurrum, 1991). The ongoing forest destruction 



poses an especially grave problem because many 
species of bats, especially endemics, depend pri- 
marily on forest (Heaney et al., 1987; Heaney and 
Utzurrum, 1991; Heideman and Heaney, 1989). 
Two species of bats (Acerodon lucifer and Dob- 
sonia chapmani) are believed to have become ex- 
tinct in the last 100 years, and many others are 
threatened (Heaney and Heideman, 1987; Heaney 
etal., 1987). 

No identification guide to the 70 species of bats 
recorded from the Philippines currently exists. The 
most recent work describing Philippine bats, Tay- 
lor's "Philippine Land Mammals", was published 
in 1934 and does not include the many species 
subsequently described or recorded from the Phil- 
ippines, nor does it reflect the many changes in 
taxonomy that have taken place over nearly 60 
years. Moreover, Taylor's keys were intended for 
use with museum study specimens, not live ani- 
mals in the field. Thus, identification of Philippine 
bats in recent years has been primarily by com- 
parison with museum specimens. This requires 
access to a comprehensive reference collection and 
is not an option for researchers in many places. 

This key is intended to permit identification of 
all bat species that have been recorded from the 
Philippines, to the extent that current knowledge 
permits. It is our hope that this key will encourage 
more research on the Philippine bat fauna, which, 
with its diversity and distribution over many is- 
lands with differing habitats and climates, serves 
as an excellent subject for studies of biogeography 
and many aspects of ecology. We also hope that a 
greater knowledge about Philippine bats will con- 
tribute to efforts toward their conservation. 



FIELDIANA: ZOOLOGY, N.S., NO. 69, OCTOBER 30, 1992, PP. 1^4 



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FIELDIANA: ZOOLOGY 



1st digit: thumb 



Claws 



Interfemoral 
membrane 




1st phalanges 

Terminal phalanges 
3rd digit 



4th digit 



5th digit 



Wing membrane 



Hind foot 
Fig. 2. Generalized pteropodid bat, with important structures indicated. 



Methods 

The descriptions and measurements in this key 
have been based primarily on examination of 
specimens in the American Museum of Natural 
History (amnh), British Museum (Natural His- 
tory) (bmnh), Delaware Museum of Natural His- 
tory (dmnh), Field Museum of Natural History 
(fmnh), Royal Ontario Museum (rom), Philippine 
National Museum (pnm), Silliman University Mu- 
seum of Natural History (su), University of Mich- 
igan, Museum of Zoology (ummz), and United 
States National Museum of Natural History 
(usnm). Our descriptions and measurements are 
based on Philippine specimens except in a few 
cases involving species that are rare in collections; 
in those cases, specimens from elsewhere or char- 
acters from published accounts were utilized. 

For standard external measurements we relied 
preferentially on data from animals that we had 
collected, but we also utilized data on specimen 
labels and measured specimens in collections. We 
took cranial measurements with dial or digital cal- 



ipers following deBlase and Martin (1974) and 
Heaney and Peterson (1984); these measurements 
are described briefly in the following section, 
Anatomy and Measurements of a Bat. 

We consulted a variety of publications as aids 
in building the keys and descriptive sections, re- 
lying heavily on Harrison (1966), Hill (1983), 
Lawrence (1939), Lekagul and McNeely (1977), 
Medway ( 1 969), Miller ( 1 907), Payne et al. ( 1 985), 
and Taylor (1934). These publications, the refer- 
ences cited in Hill (1983), and publications re- 
ferred to in the sections for each bat family should 
be consulted for descriptions of characters not 
treated in the key. 

Anatomy and Measurements 
of a Bat 

Familiarity with basic terminology on the anat- 
omy of a bat is necessary to use this key; Figure 
2 shows the most important external structures. 
The external ear (pinna) has two structures that 
can be useful for identification. The tragus is a 



Fig. 1 . Map of the Philippines. Recent islands are outlined by continuous lines, and the extent of late Pleistocene 
islands is indicated by the shaded areas. 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 




Fig. 3. Left external ears (pinnae) of representative 
bats. A, Pteropodidae; B, Rhinolophidae; C, Vespertil- 
ionidae. 



projection from the base of the ear (fig. 3C); this 
structure is absent in members of the Pteropodidae 
(fig. 3 A) and Rhinolophidae (fig. 3B) and is quite 
small in the Molossidae. The antitragus is the flap 
along the lower posterior margin of the ear (figs. 
3B, 12); it is especially well developed in rhinol- 
ophids and most molossids. Rhinolophids and 
megadermatids have noseleaves, structures formed 
from elaborate folds of skin on their noses (figs. 
8B,C, 12); bats of other families have no such 
structure. 

All bats have four types of teeth: incisors, ca- 
nines, premolars, and molars (fig. 5). All bats pos- 
sess a pair of large, sharp, conical teeth called ca- 
nines on the upper and lower jaws. The smaller 
teeth at the front of the mouth, anterior to the 



canines, are incisors. Posterior to the canines are 
the premolars and molars, collectively called cheek 
teeth. The large teeth in this series are referred to 
as molariform teeth. Molariform teeth are usually 
low and broad with cutting or grinding surfaces; 
the anterior cheek teeth are tiny pegs in some spe- 
cies. 

External and cranial measurements provide 
substantial aid in identification; we have therefore 
included tables containing typical adult measure- 
ments. The definitions of standard external mea- 
surements are illustrated in Figure 4. The cranial 
measurements included in the tables are shown in 
Figure 5 and are defined as follows: condylobasal 
length (CBL), distance from the posterior edge of 
the occipital condyles to the anterior tip of the 
premaxillaries; condylocanine length (CCL), dis- 
tance from the posterior edge of the occipital con- 
dyles to the anterior edge of the base (alveolus) of 
the canines; and maxillary toothrow (C to last M), 
distance from the posterior edge of the last upper 
molar to the anterior edge of the upper canine, 
taken along the bone line of the alveolus. 

Use of the Key 

Using the key most effectively requires several 
steps that are explained in detail in the following 
sections. First, the bat should be identified as to 



Total length 



Ear 




f Oni I 



Fig. 4. Bat with external measurements indicated. 



FIELDIANA: ZOOLOGY 



Incisors 




.Condylobasal length. 
.Condylocanine length. 




Fig. 5. Bat skull with teeth and cranial measurements indicated. 



age and sex. It should then be identified to family 
with the Key to Families, and then to species with 
the key to that family. Next, all external mea- 
surements should be taken and matched against 
those in the tables of measurements; if the mea- 
surements do not match, the identification is ques- 
tionable and the process should be begun again 
from the first step. Finally, if a skull is available, 
it should be visually compared with Figures I9- 
60 to see if it matches the features of that species 
or genus. 



Determination of Age and Sex 

The age of bats can be estimated by the degree 
of ossification of the joints in the digits of the wing 
(Anthony, 1988). In juvenile bats, the joints have 
cartilaginous discs where growth takes place. When 
a light is shown through the wing of a live bat, the 
bands of cartilage at the joints appear partly trans- 
lucent (fig. 6B). In adult bats, the bones are fully 
ossified and the joints appear opaque (fig. 6 A). The 
shape of the joints of the digits also differs between 
juvenile and adult bats; this is especially useful in 
determining the age of animals prepared as stuffed 
skins or fluid-preserved specimens. In juvenile bats 



the joints are swollen and tapered (fig. 6B), whereas 
in adult bats the joints are knobby and more dis- 
tinct from the bone shaft (fig. 6A). 

Bats can be sexed by examination of the external 
genitalia. Males have a conspicuous penis (except 



B 



Fig. 6. Joints in digits of wing. Stippled areas rep- 
resent bone and open areas represent cartilage. A, adult; 
B, juvenile. 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



subadult male Rousettus, in which the penis is 
retracted into the abdomen). Both sexes possess 
axillary nipples on the upper chest, usually near 
the armpit (axilla). However, the nipples of adult 
females are more prominent than those of males. 
In female rhinolophids and megadermatids, a pair 
of inguinal papillae that look like nipples is present 
just anterior to the genitals. 



Identification to 
Family and Species 



when overlap in the ranges of two or more species 
means that the measurement cannot be used to 
fully distinguish between the species. When other 
measurements are particularly useful for separat- 
ing similar species, they are also given in the key. 
Measurements should not be used to identify ju- 
venile bats, although hind foot length is sometimes 
useful because the feet of juvenile bats approach 
adult size long before most other structures have 
ended growth. 

Comparison with Skull Drawings 



Following determination of age and sex, the bat 
should be identified to family with the Key to 
Families, and then to species with the key for the 
appropriate family. The keys consist of sets of al- 
ternatives arranged in order of increasing restric- 
tiveness. Alternatives belonging to the same set 
are preceded by the same number with different 
superscripts (e.g., 1,1', 1"). For each set, the user 
should choose the best alternative; this indicates 
either the next set of alternatives to be considered, 
or the name of the family or species. All distin- 
guishing characters should be checked carefully 
before a choice is made. This will minimize the 
likelihood of incorrectly identifying a bat. Re- 
searchers should bear in mind that this key in- 
cludes only those species now known from the 
Philippines, and that discovery of undescribed 
species and previously unreported species is likely. 

Figures illustrating important descriptive char- 
acteristics are referred to in the key and should be 
consulted. Fur coloration can be useful in identi- 
fication; it should be observed on a dry animal or 
study skin. 



Use of Measurement Tables 

Once a bat has been identified to species, it is 
good practice to check if its measurements agree 
with those given in the tables. Cranial measure- 
ments should be taken with calipers. External 
measurements can be taken with a ruler, although 
calipers provide more accurate measurements of 
forearm length and of small structures such as parts 
of the noseleaves of rhinolophids. 

Because forearm length is a very useful mea- 
surement for identifying adult bats, ranges for fore- 
arm length are given in the keys for each species, 
usually at the end of the final description for that 
species. Note that forearm lengths are given even 



Skulls are very important in identification of 
bats. Thus, although we have written this key with 
the intention that live animals and preserved spec- 
imens can be identified without examination of 
their skulls, we have included skull drawings of 
common Philippine bats (figs. 1 9-60). The draw- 
ings will aid in identification of museum speci- 
mens with cleaned skulls and will provide a means 
of verifying an identification based on external 
characteristics. The skull drawings also serve as a 
guide to head shape. Even if use of the key yields 
an apparently unambiguous identification, it is al- 
ways best if a skull can be examined from a vouch- 
er specimen. The skull should be compared with 
the figures, and cranial measurements should be 
taken and compared with those in the tables. Den- 
tition, although most easily visible on a skull, can 
also be observed, at least partially, on a live animal 
and should be examined when possible. The mouth 
can be gently pried and held open by a toothpick, 
and the lips can be lifted to view the cheek teeth. 
With small bats a magnifying lens is a great help. 
Note that if the jaws are closed, some teeth, par- 
ticularly the canines, may obstruct the view of 
others. 

Users of this key should not overlook the fact 
that variation in both quantitative and qualitative 
characters is present in all species. Whenever pos- 
sible, samples of more than one individual for each 
species should be examined and the most common 
character states used in the key. 

The identities of a few species of Philippine bats 
that are extremely poorly represented in collec- 
tions (sometimes only by the holotype) are uncer- 
tain. When possible, these species have been in- 
cluded in the key on the basis of whatever 
descriptions are available, and further comments 
on their identification are provided in the Notes 
section. We also comment briefly on two new re- 
cords of bats for the Philippines in this section. 



FIELDIANA: ZOOLOGY 



Collecting and Preserving 
Voucher Specimens 

In any study, several individuals of each species 
should be preserved as voucher specimens and 
deposited in a museum collection. Voucher spec- 
imens are necessary to verify identification, since 
many species can be identified definitively only 
after careful study of the external and cranial mor- 
phology. Such specimens are invaluable for adding 
to our current, very limited knowledge of Philip- 
pine bats, particularly of distribution and inter- 
and intraspecific variation. Specimens should be 
collected and prepared carefully, following all reg- 
ulations for the scientific study of wildlife. 

Bats are most easily preserved in fluid. They 
should be killed in a quick and painless manner, 
examined for reproductive condition, and weighed, 
and their standard external measurements taken 
(total length, tail length, ear length, and hind foot 
length). The date, habitat, collection locality, el- 
evation, collector, collector's field number, and 



external measurements should be recorded in In- 
dia ink (which is best) or pencil (which is usually 
acceptable), not with other kinds of ink (which are 
not permanent). These data should accompany the 
specimen as part of the permanent record. The 
specimen should have a permanent label attached 
that bears the collector's name and field number; 
other data may go onto the same label or into a 
field catalog. The specimens should then be rinsed 
lightly with soapy water, injected with a 10% so- 
lution of formalin, and immersed in a 1 0% solu- 
tion of formalin. After about three days, they can 
be rinsed with clean water and transferred to 70% 
alcohol (methanol is best) for permanent storage. 
Skulls can be extracted and cleaned in the muse- 
um. 

The Philippine National Museum, University 
of the Philippines at Los Banos, Silliman Univer- 
sity, and a number of other institutions maintain 
research collections and accept voucher specimens 
for their permanent collections, and the staff may 
be able to help with identifications. 



Key to Families of Philippine Bats 

1 . Interfemoral membrane is absent or reduced, forming narrow margin along insides of legs; tail short 
(up to 20% of body length) or absent, never completely enclosed by interfemoral membrane (fig. 7 A); 
both thumb and second finger with claw (except Eonycteris and Dobsonia; fig. 2); both tragus and 
antitragus absent (figs. 3A, 8A); ear margin forms continuous ring . . . Megachiroptera: Pteropodidae 

1'. Interfemoral membrane is a continuous expanse of skin stretching between legs (figs. 7B-F; except 
Coelops, which has a concave posterior margin to the membrane and no tail); tail present (except 
Megaderma and Coelops, which both possess noseleaves, structures not present in any pteropodids), 
usually comprising more than 20% of body length; the second finger does not have a claw; either 
tragus (fig. 3C) or antitragus (fig. 3B) or both present; ear margin does not form continuous ring . . 
(Microchiroptera) 2 

2. Noseleaf present (figs. 8B,C) 3 

2'. Noseleaf absent (figs. 8D-F) 4 

3. Large ears connected at top of forehead (fig. 8B); tragus long and forked; external tail absent but 
interfemoral membrane well developed (fig. 7B) Megadermatidae 

3'. Ears not connected across top of forehead; tragus absent, but antitragus usually well developed (fig. 
3B); tail present, enclosed by interfemoral membrane except at extreme tip (fig. 7D; except Coelops, 
which has no tail and a reduced interfemoral membrane Rhinolophidae 

4. Tail extends to posterior margin of interfemoral membrane (extreme tip may project 1-2 mm beyond 
membrane; fig. 7C); ears variable, usually not fleshy (fig. 8F); anterior edge of bony palate deeply 
emarginate Vespertilionidae 

4'. Tail emerges dorsally from interfemoral membrane but is shorter than membrane when legs and 
membrane are outstretched (fig. 7F); ears not noticeably thick and fleshy (fig. 8D); anterior edge of 
bony palate deeply emarginate Emballonuridae 

4*. Tail projects beyond posterior margin of interfemoral membrane for over half its length (fig. 7E); 

ears thick and fleshy (fig. 8E); anterior edge of bony palate continuous, not emarginate 

Molossidae 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 







mm 




Fig. 7. Tails of representatives of Philippine bat families. A, Pteropodidae (Rousettus); B, Megadermatidae 
(Megaderma); C, Vespertilionidae (Miniopterus); D, Rhinolophidae (Rhinolophus); E, Molossidae (Chaerephon); F, 
Emballonuridae (Taphozous). 



Family Pteropodidae: 
Fruit Bats 

All fruit bats are characterized by dog-like heads 
(fig. 8A), with eyes that are proportionately larger 



than those of most other bats; the large eyes are 
associated with their dependence on vision for ori- 
entation and their lack of echolocation (sonar) sys- 
tems (except for the very simple and limited one 
used by Rousettus). They do not have the elaborate 



Fig. 8. Heads of representatives of Philippine bat families (not to same scale). A. Pteropodidae (Rousettus). Note 
that the ear margin is continuous, neither tragus nor noseleaf is present, the eyes are large, and the face is dog-like. 
B. Megadermatidae (Megadermd). Note that a noseleaf is present, the tragus is forked, and the large ears are connected 
across the top of the forehead. C. Rhinolophidae (Rhinolophus). Note the elaborate noseleaf, the well-developed 



FIELDIANA: ZOOLOGY 




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antitragus, and the absence of a tragus. D. Emballonuridae (Taphozous). Note that there is no noseleaf and that a 
tragus is present. E. Molossidae (Chaerephon). Note that there is no noseleaf and that the ears are thick and fleshy. 
F. Vespertilionidae (Miniopterns). Note that there is no noseleaf and that a tragus is present. 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



folds of skin in the nasal region (also associated 
with the use of echolocation) that are present in 
two other families of bats in the Philippines 
(Megadermatidae and Rhinolophidae). The exter- 
nal portion of the ear is simple and of moderate 
size, and the margin of the ear forms a continuous 
ring. Fruit bats do not have a tragus or antitragus 
(figs. 3A, 8A). All but three species (Dobsonia 
chapmani, Eonycteris robusta, and E. spelaea) have 
a claw on the second digit (fig. 2), along the leading 
edge of the wing; bats belonging to other families 
lack this claw. Many species have no tail, and in 
none does the tail comprise more than 20% of the 
body length. Their size varies greatly, with forearm 
lengths ranging from 40 to 215 mm, and weight 
from about 16 g (Alionycteris paucidentata) to at 
least 1,200 g (Acerodon jubatus; table 1). 

Most species show substantial sexual dimor- 
phism; males are larger than females and often 
have broader, more heavily muscled heads. Males 
of many species possess a ruff of fur (often colored 
yellow or rusty red) around the neck and on the 
shoulders that is lacking or poorly developed in 
females. Subadults of both sexes resemble adult 
females. Parous adult females of all species have 
a pair of axillary nipples that are large and prom- 
inent; males and subadult females have very small 
but visible nipples. 

The skulls of all pteropodids are characterized 
by the presence of prominent postorbital process- 



es, and all but a few species have postorbital fo- 
ramina (figs. 19-32). The dental formula is vari- 
able. Most species have broad, blunt, rounded 
molariform teeth, but the nectarivorous species 
have greatly reduced premolars and molars. All 
species have long, prominent canines. In many 
species, males have longer canines than do females 
and often have more strongly developed cranial 
crests. 

As the English name for the family implies, most 
fruit bats feed on fruit, but three species {Eonyc- 
teris robusta, E. spelaea, and Macroglossus mini- 
mus) feed primarily on nectar and pollen. Fruit 
bats often are abundant, being uncommon only in 
upper montane and mossy forest (Heaney and 
Rickart, 1990; Heaney et al., 1989). In forested 
areas, small to medium-sized species roost in hol- 
low trees and in foliage, either alone or in groups; 
the large flying foxes roost in exposed treetops. 
Several species roost in caves, often forming col- 
onies in the hundreds or thousands. 

Andersen (1912) provided descriptions of most 
species of pteropodids found in the Philippines. 
Additional descriptions may be found in Berg- 
mans (1975, 1978), Francis (1989), Heaney and 
Peterson (1984), Klingener and Creighton (1984), 
Kock (1969a,b,c), Musser et al. (1982), Peterson 
and Fenton (1970), Rookmaaker and Bergmans 
(1981), and Yoshiyuki (1979). 



Key to Pteropodidae 






1 . Claw on thumb but not on second digit 2 

1 '. Claws on both thumb and second digit (fig. 2) 4 

2. Wings attach to body along midline of back; two upper incisors and two minute lower incisors; 
forearm 123-131 mm; skull as in Figure 21 Dobsonia chapmani 

2'. Wings attach along sides of body; four upper and four lower incisors, all very small; forearm 67- 
82 mm (Eonycteris) 3 

3. Pair of prominent 2-6-mm-long kidney-shaped glands lateral to anus; tail 12-20 mm; forearm 67- 
80 mm; skull as in Figure 23 Eonycteris spelaea 

3'. No glands near anus; tail 20-28 mm; forearm 67-82 mm; skull as in Figure 22 

Eonycteris robusta 

4. Forearm 94-2 1 5 mm; tail absent; four upper and four lower incisors 5 

4'. Forearm 41-92 mm; tail present or absent; number of incisors varies between species 13 

5. Wings with prominent pale blotches, particularly along thumb and anterior edge of wing adjacent 
to first and second digit, especially at wing tips; forearm 135-141 mm Pteropus leucopterus 

5'. Wings dark brown without prominent pale blotches; forearm 94-21 5 mm 6 

6. Pelage on dorsal surface of lower back pale brown or gray 7 

6'. Pelage on dorsal surface of lower back dark brown or black, sometimes with yellow flecks .... 8 

7. Forearm 132-165 mm; condylobasal length 65-70 mm; three cusps on second and third upper 
molariform teeth, including a well-developed anterolingual cusp Acerodon leucotis 

7'. Forearm 94-1 13 mm; condylobasal length 46-52 mm; two cusps on second and third upper mo- 
lariform teeth (no anterolingual cusps); skull as in Figure 30 Pteropus pumilus 



10 



FIELDIANA: ZOOLOGY 






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INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



11 






D 




Fig. 9. Frontal views of fruit bat skulls, showing incisors and canines (not to same scale). A, Otopteropus carti- 
lagonodus (Alionycteris paucidentata and Haplonycteris fischeri have an equal number of incisors); B, Cynopterus 
brachyotis; C, Nyctimene rabori; D, Ptenochirus minor {Ptenochirus jagori incisors are very similar). 



8. Forearm 165-215 mm; condylobasal length 68-86 mm; fur on upper back sometimes completely 
black 9 

8'. Forearm 1 1 8-1 52 mm; condylobasal length 54-66 mm; fur on upper back never completely black, 
usually golden 11 

9. Dorsal pelage may be completely dark brown or black; if upper back is golden, the posterior margin 
of the golden area forms a sharply defined transverse line with the dark brown lower back; tips of 
ears nearly pointed; two cusps on second and third upper molariform teeth (no anterolingual cusps); 
condylobasal length 68-79 mm; forearm 1 79-204 mm; skull as in Figure 31 .... Pteropus vampyrus 

9'. Dorsal pelage never completely dark brown or black; a golden patch on top of head, extending 
anterior to line between ears, is always present; the golden dorsal pelage never forms a transverse 
line along the edge of the dark brown of the lower back; tips of ears bluntly rounded; three cusps 
on second and third upper molariform teeth, including a well-developed anterolingual cusp; forearm 
165-215 mm; condylobasal length 71-86 mm 10 

10. Forearm 167-170 mm; condylobasal length 71-74 mm (occurs only on Panay Island; see Notes) 

Acerodon lucifer 

10'. Forearm 165-215 mm; condylobasal length 72-86 mm; skull as in Figure 19 ... Acerodon jubatus 

11. Forearm 1 18-133 mm; condylobasal length 53-62 mm (occurs only from Zamboanga to Sulu) . . 

Pteropus speciosus 

11 '. Forearm 133-152 mm; condylobasal length 56-66 mm 12 

12. Lower legs heavily furred nearly to ankle; forearm 133-152 mm (see Notes) 

Pteropus dasymallus 

12'. Lower legs nearly naked; forearm 136-149 mm; skull as in Figure 29 ... Pteropus hypomelanus 

1 3. Tail absent 14 

1 3'. Tail present (fig. 7A; it may be small, so look carefully) 19 

14. Two small lumps of soft white tissue on each ear, at anterior and posterior margins; forearm 43- 
50 mm Otopteropus cartilagonodus 

14'. No lumps of soft white tissue at margins of ears; forearm 41-91 mm 15 

15. Forearm 80-91 mm; well-developed secondary cusps on canines (one on upper canines, two on 
lower canines); six cusps on last two upper and lower molariform teeth; dorsal surface of hind feet 
thickly furred; skull as in Figure 25 Harpyionycteris whiteheadi 

15'. Forearm 41-53 mm; canines do not have well-developed secondary cusps; two or three cusps on 
molariform teeth; dorsal surface of hind feet may or may not be thickly furred 16 

1 6. Muzzle long and slender (fig. 1 0A); nostrils are not tubular and do not project beyond rest of muzzle; 
wing membrane attaches on top of foot, above gap between third and fourth toes from outside (fig. 



12 



FIELDIANA: ZOOLOGY 








Fig. 10. Dorsal views of fruit bat muzzles (not to same scale). A, Macroglossus; B, Haplonycteris; C, Nyctimene; 
D, Rousettus; E, Cynopterus. 



1 1A); teeth, except canines, greatly reduced; forearm 41-45 mm; skull as in Figure 26 

Macroglossus minimus 

16'. Muzzle short and broad; nostrils tubular (as in fig. 10B); wing membrane attaches on side of foot 
(on or above outermost toe; fig. 1 1C); teeth robust; forearm 43-53 mm 17 

17. Four upper and two lower incisors (fig. 9D); forearm 45-52 mm Megaerops wetmorei 

17'. Two upper and two lower incisors (as in fig. 9 A); forearm 43-53 mm 18 

18. Band of pale fur along dorsal surface of forearm; interfemoral membrane present; forearm 46-53 
mm; skull as in Figure 24 Haplonycteris fischeri 

18'. No band of pale fur along dorsal surface of forearm; no interfemoral membrane; forearm 45-50 
mm Alionycteris paucidentata 

19. Ears and skin on dorsal surface of bones of wings with prominent pale yellow spots; dark stripe 
along most of dorsal midline; two upper and no lower incisors (fig. 9Q; nostrils elongated into tubes 
about 2-3 mm long (fig. IOC); forearm 71-79 mm; skull as in Figure 27 Nyctimene rabori 

19'. Ears and wings without yellow spots; no stripe along dorsal midline; four upper incisors and at 
least two lower incisors; nostrils not elongated into tubes (although when viewed from above they 
may appear slightly tubular); forearm 45-92 mm 20 

20. Muzzle moderately long and tapered (fig. 10D); anterior surface of upper canines with vertical 
groove (not always prominent); wing membrane terminates above gap between outermost toe and 
second toe from outside (fig. 1 1 B); forearm 80-92 mm; skull as in Figure 32 

Rousettus amplexicaudatus 

20'. Muzzle short and broad (fig. 10E); anterior surface of upper canines smooth, not grooved; wing 
membrane terminates either on outermost toe (fig. 1 1C) or above gap between outermost toe and 
second toe from outside (fig. 1 1 B); forearm 45-92 mm 21 

21. Four upper and two lower incisors (outer pair of upper incisors much smaller than inner pair; fig. 
9D) 22 

21'. Four upper and four lower incisors (fig. 9B) 23 

22. Forearm 76-90 mm; skull as in Figure 28 Ptenochirus jagori 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



13 



22'. Forearm 60-78 mm (Greater Mindanao only) Ptenochirus minor 

22". Forearm 45-52 mm Megaerops wetmorei 

23. Anterior edges of ears pale; wing membrane terminates on side of foot, on outermost toe (fig. 1 1C); 

cheek teeth moderate in size; forearm 58-68 mm; skull as in Figure 20; (see Notes) 

Cynopterus brachyotis 

23'. No pale rims to ears; wing membrane terminates above gap between first and second toe from 

outside (fig. 1 1 B); cheek teeth greatly enlarged, squarish in outline with large cusps and ridges; 

forearm about 92 mm Dyacopterus spadiceus 



Family Emballonuridae: 
Sheath-Tailed Bats 

Sheath-tailed bats are small to moderate-sized 
bats (forearm 44-71 mm; table 2). They are the 
only bats in the Philippines in which the tail per- 
forates the dorsal surface of the interfemoral mem- 
brane (fig. 7F), so that the basal portion (usually 
about half) of the tail is enclosed within the mem- 
brane and the distal portion is free, lying on the 
dorsal surface of the membrane. The eyes are small 
to moderate in size, and the ears are simple and 
of moderate size (fig. 8D). A tragus is always pres- 
ent. The muzzle is rather pointed, with the nostrils 
at the tip; it does not have a noseleaf. 



The skulls of emballonurids have postorbital 
processes that are very well developed (figs. 33- 
35); these processes are absent in vespertilionids, 
which are superficially the most similar to em- 
ballonurids. Emballonurid premaxillaries are small 
and delicate; they do not fuse to each other at the 
anterior midline and are attached to the maxil- 
laries by flexible connective tissue. The anterior 
edge of the palate has a deep emargination. 

All of the sheath-tailed bats in the Philippines 
feed on insects. Common roosting sites are caves, 
hollow trees, and attics of buildings; they usually 
aggregate in moderate to large groups (ten to sev- 
eral hundred). 



Key to Emballonuridae 

1. Two distinct pairs of upper incisors; forearm 44-49 mm; skull as in Figure 33 

Emballonura alecto 

1 '. One pair of upper incisors; forearm 61-71 mm 2 

2. Forearm 6 1-65 mm; dorsal pelage pale brown to sandy at tips, very pale at bases; skull as in Figure 
35 Taphozous melanopogon 

2'. Forearm 66-71 mm; dorsal pelage very dark brown, usually with flecks of white; dark hairs do not 
have pale bases; skull as in Figure 34 Saccolaimus saccolaimus 



Family Megadermatidae: 
False Vampire Bats 



This is the least diverse of bat families in the 
Philippines, with only a single species present, 
Megaderma spasma (fig. 8B). M. spasma is a me- 
dium-sized bat (forearm 57-63 mm; table 2) easily 
recognized by its large ears that meet across the 
top of the forehead, erect, simple noseleaf, and 
lack of tail (although the interfemoral membrane 
is well developed). 

The skull of Megaderma spasma is of moderate 
size and its stoutly constructed (fig. 36). The pre- 
maxillaries are absent, so there are no upper in- 



cisors, and the anterior edge of the palate has a 
deep emargination. The postorbital processes are 
very poorly developed. The upper canines have a 
large secondary cusp. 




Fig. 1 1 . Posterior attachment of wing membrane on 
left foot. A, Macroglossus; B, Rousettus; C, Ptenochirus/ 
Haplonycteris. 



14 



FIELDIANA: ZOOLOGY 



Table 2. Measurement ranges of at least 10 individuals of adult Philippine Emballonuridae and Megadermatidae. 
Measurements, as denned in text, were taken from Philippine specimens. 



Species 



CBL 



CCL 



C'-last 
M 



Total 
length 



Tail 



Hind 
Foot 



Ear 



Fore- 
arm Weight 



Embailonura alecto 13.3-14.2 12.5-13.4 5.0-5.8 56-69 9-12 7-10 12-16 44-49 4-6.5 

Saccolaimus saccolaimus 20.9-23.3 8.8-9.9 103-117 20-28 15-19 16-20 66-71 28-36 

Taphozous melanopogon 19.6-20.6 8.1-8.8 100-111 20-25 11-15 20-24 61-65 20-29 

Megaderma spasma - 21.2-23.6 8.7-9.4 70-89 18-22 36-43 57-63 21-27 



Megaderma spasma feeds primarily on large in- 
sects (especially cicadas and katydids) and occa- 
sionally on small vertebrates such as lizards, frogs, 
and small birds (D. Balete, pers. comm.). It has 
been found roosting in caves and in hollow trees, 
singly and in small groups. 

Because only a single species is present, no key 
to the family is necessary. 

Family Rhinolophidae: Horseshoe 
and Leaf-Nosed Bats 

The family Rhinolophidae is composed of about 
ten genera and over 100 species; it is represented 
in the Philippines by three genera (Coelops, Hip- 
posideros, and Rhinolophus) and at least 1 7 species 
(see Notes). All rhinolophids have a noseleaf, a 
structure consisting of elaborate folds of skin in 
the nasal region. The shape of the noseleaf varies 
among species and is an important character for 
identification (fig. 12). In members of the genus 
Rhinolophus, the posterior noseleaf is long and 
pointed (fig. 12A) and the anterior noseleaf is 
horseshoe-shaped. Between the anterior and pos- 
terior noseleaves lies the sella, an anterior-facing 
structure that is connected to the posterior noseleaf 
by the connecting process. In some species, sup- 
plementary leaflets are present lateral and ventral 
to the posterior noseleaf (fig. 12A). In Coelops and 
Hipposideros, the posterior noseleaf is low and 
rounded (fig. 1 2B) and may be divided into pock- 
ets by vertical septa. The intermediate noseleaf is 
a cushion-like structure. Supplementary leaflets are 
sometimes present lateral and ventral to the an- 



terior noseleaf (not shown in fig. 12B, but see fig. 
12A). Between the nostrils is the internarial sep- 
tum. Rhinolophids lack a tragus, but many spe- 
cies, especially those belonging to the genus Rhi- 
nolophus, possess a well-developed antitragus, a 
flap of skin on the lower posterior margin of the 
ear (fig. 1 2). Most rhinolophids are small (forearm 
34-60 mm), but a few are medium-sized (forearm 
up to 90 mm; table 3). 

The skulls of most rhinolophids are delicate and 
slender, although some, especially those of the 
larger Hipposideros, are fairly stout. Postorbital 
processes are absent. Premaxillary bones are pres- 
ent but are small and connected to the skull by 
cartilaginous articulations, and so are movable in 
live or freshly killed animals. Incisors are present 
but are moderate to very small. The lower incisors 
are trilobed, although this is often inconspicuous. 
Many species have a strongly expanded nasal re- 
gion (which supports the noseleaf structures), giv- 
ing the skull a strongly sinuous dorsal profile in 
lateral view (figs. 37-46). 

Rhinolophids are insectivorous. Some species 
roost in caves, in numbers ranging from a few to 
several hundred. Other species roost in hollow 
trees, hollow fallen logs, and other sites in forest. 
Although rhinolophids usually comprise a small 
proportion of captures in mist nets, at times a 
single species may be the most common bat spe- 
cies netted at a given site. 

For further descriptions of members of the ge- 
nus Hipposideros, see Hill ( 1 963a) and Jenkins and 
Hill (1981). For Rhinolophus, see Andersen 
(1905a,b,c,d). 



Key to Rhinolophidae 

1 . Posterior noseleaf pointed (fig. 1 2 A); six pairs of lower cheek teeth (Rhinolophus) 2 

1'. Posterior noseleaf low and flattened (fig. 12B); five pairs of lower cheek teeth 8 

2. Forearm 68-73 mm Rhinolophus rufus 

2'. Forearm 38-57 mm 3 

3. Dorsal tip of connecting process sharply pointed (fig. 13A); forearm 47-49 mm 

Rhinolophus acuminatus 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



15 



Posterior 
noseleaf 




^^X, 




Fig. 12. Noseleaves of Rhinolophus and Hipposideros. A, Rhinolophus; B, Hipposideros. 

3'. Dorsal tip of connecting process not pointed (figs. 13B,C); forearm 38-57 mm 4 

4. Antitragus nearly triangular (as in fig. 12 A); cup at base of sella (immediately above nostrils) 8-10 
mm wide, almost as wide as anterior noseleaf; forearm 51-57 mm; skull as in Figure 45 

Rhinolophus philippinensis 

4'. Antitragus most nearly rectangular; cup at base of sella 2-4 mm wide, no more than half as wide 
as anterior noseleaf; forearm 38-56 mm 5 

5. Connecting process attaches below tip of sella (fig. 1 3B) 6 

5'. Connecting process attaches at tip of sella (fig. 1 3C) 7 

6. Ears 25-26 mm; sella 3-4 mm wide; forearm 43-44 mm; skull as in Figure 44 (see Notes) 

Rhinolophus macrotis 

6'. Ears 17-21 mm; sella 1-2 mm wide; forearm 38-44 mm; skull as in Figure 46 

Rhinolophus virgo 

7. Forearm 43-46 mm; condylocanine length 16.2-17.4 mm; maxillary toothrow 6.7-7.5 mm; skull 
as in Figure 42 (see Notes) Rhinolophus arcuatus-s 



16 



FIELDIANA: ZOOLOGY 



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INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



17 



7'. Forearm 47-50 mm; condylocanine length 17.9-19.0 mm; maxillary toothrow 7.7-8.1 mm; skull 
as in Figure 41 (see Notes) Rhinolophus arcuatus-l 

7". Forearm 49-55 mm; condylocanine length 19.9-21.0 mm; maxillary toothrow 8.3-9.2 mm; skull 
as in Figure 43 (see Notes) Rhinolophus inops 

1'" . Forearm 51-56 mm; condylocanine length about 21.8 mm; maxillary toothrow 9.0-9.5 mm (see 
Notes) Rhinolophus subrufus 

8. Tail absent; anterior noseleaf bilobed with two lateral leaflets underneath that project downward; 
no ridges on ears; well-developed secondary cusps on upper canines; forearm about 34 mm .... 

Coelops hirsuta 

8'. Tail present, greater than 15 mm; anterior noseleaf undivided, without leaflets that project down- 
ward; ridges on ears; only one cusp on upper canines; forearm 37-89 mm .... (Hippos ideros) 9 

9. Forearm 77-89 mm; cream-colored patch of fur anterior to leading edge of wing; skull as in Figure 
38 Hipposideros diadema 

9'. Forearm 37-50 mm 10 

10. Two pairs of supplementary leaflets (see fig. 12 A) lateral to anterior noseleaf 11 

10'. No supplementary leaflets lateral to anterior noseleaf 12 

11. Inner pair of lateral leaflets meet under anterior noseleaf but outer pair do not; forearm 37-40 mm; 
skull as in Figure 40 Hipposideros pygmaeus 

11'. Both pairs of lateral leaflets do not meet under anterior noseleaf; forearm 44-50 mm 

Hipposideros cervinus 

12. No vertical septa in posterior noseleaf and tail >30 mm; forearm about 47 mm (see Notes) .... 

Hipposideros coronatus 

12'. Vertical septa may or may not be present in posterior noseleaf; if they are absent, then tail 18-24 
mm; forearm 38-48 mm 13 

13. Anterior noseleaf 5.5-7.0 mm wide, intermediate noseleaf 5.0-6.0 mm wide, posterior noseleaf 
6.0-8.0 mm wide; hind foot length 10-12 mm; forearm 42-48 mm; skull as in Figure 39 

Hipposideros obscurus 

13'. Anterior noseleaf 4.0-5.0 mm wide, intermediate noseleaf 3.5^4.0 mm wide, posterior noseleaf 
4.5-5.5 mm wide; hind foot length 7-10 mm; forearm 38-43 mm 14 

14. Internarial septum not swollen at base; forearm about 42 mm Hipposideros bicolor 

14'. Internarial septum swollen at base; forearm 38-43 mm; skull as in Figure 37 ... Hipposideros ater 



Family Vespertilionidae: 
Evening Bats 

This is an exceptionally large and diverse family, 
with about 40 genera and 275 species worldwide 
and about 1 1 genera and 22 species in the Phil- 
ippines. Vespertilionids are identified as having a 
long tail that is completely enclosed by the inter- 
femoral membrane (except occasionally the ex- 



B 






Fig. 13. Lateral views of connecting processes of some 
rhinolophids. A, R. acuminatum; B, R. macrotis (that of 
R. virgo is very similar); C, R. subrufus/ R. arcuatus. 



treme tip; fig. 7C), a simple face without a noseleaf 
(among species in the Philippines; fig. 8F), and a 
well-developed tragus. Most vespertilionids are 
small to medium-sized (forearm 22-55 mm; table 
4) and have small eyes. 

The skulls of vespertilionids are generally small, 
with a great variety of shapes and degrees of ro- 
bustness (figs. 47-59). The braincase is high and 
domed in some genera (e.g., Miniopterus, Myotis; 
figs. 48-50, 52-55) and exceptionally low and flat- 
tened in others (e.g., Tylonycteris). They all lack 
postorbital processes. All species have a deep 
emargination at the anterior end of the palate, so 
that the incisors on either premaxillary are widely 
separated. The molariform teeth are generally typ- 
ical of those of insectivorous bats, with broad 
crowns and sharp, shearing crests. 

All of the vespertilionids in the Philippines are 
insectivorous. Roosting sites include caves, build- 
ings, foliage, hollow trees, unfurled banana leaves, 



18 



FIELDIANA: ZOOLOGY 






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INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



19 



and bamboo internodes. Some species have been Many vespertilionid species that occur in the 

captured only within forest, whereas others are Philippines are described in Hill (1983). Other 

common in urban areas. Species that roost in caves useful descriptions can be found in Francis and 

and buildings often form large colonies, ranging Hill (1986), Heaney and Alcala( 1986), Hill (1963b, 

from hundreds to tens of thousands. 1965, 1966, 1971), Kock (1981), and Tate (1942). 



Key to Vespertilionidae 

Nostrils elongated into short tubes that open laterally (fig. 1 4 A); long orange hairs on dorsal surface 
of interfemoral membrane and feet; tragus straight and pointed, at least half length of ear (fig. 1 5 A) 

2 

Nostrils not elongated into tubes (fig. 1 4B); hairs on interfemoral membrane very short and almost 

invisible; tragus shape and size varies between species 3 

Forearm 36-39 mm; five well-developed pairs of upper cheek teeth; skull as in Figure 51 

Murina cyclotis 

Forearm about 5 1 mm; fifth pair of upper cheek teeth minute or absent (see Notes) 

Harpiocephalus harpia 

Posterior margin of ear has a pronounced concave inflection near tip and forms a large, rounded 
flap below inflection, producing funnel-shaped ears (fig. 15B); tragus straight and pointed, at least 

half length of ear 4 

No large, rounded flap on posterior margin of ear (figs. 1 5C,D); tragus shape and size varies between 

species 7 

Longitudinal groove on anterior surface of upper canines; individual hairs of dorsal pelage with 
four bands of color (dark at base, then buff, then dark brown, tip buff or golden), producing a flecked 

or salt-and-pepper appearance; forearm about 39 mm Phoniscus jagorii 

Upper canines not grooved; hairs of dorsal pelage have two or three bands of color; forearm 30- 

35 mm {Kerivoula) 5 

Bases of dorsal hairs pale; forearm about 35 mm Kerivoula pellucida 

Bases of dorsal hairs dark; forearm 30-35 mm 6 

Anterior two pairs of upper and lower premolars narrow and elongate, oval in cross-section; forearm 

about 30 mm Kerivoula whiteheadi 

Anterior two pairs of upper and lower premolars approximately round in cross-section; forearm 

34-35 mm; skull as in Figure 47 Kerivoula hardwickii 

One pair of upper incisors, which are large and conical; dorsal pelage rusty orange; forearm 47-53 

mm; skull as in Figure 59 Scotophilus kuhlii 

Two pairs of upper incisors; color of dorsal pelage varies between species; forearm 22-55 mm ... 8 

Tragus erect and tapering (fig. 1 5C) {Myotis) 9 

Tragus blunt (e.g., fig. 1 5D) 12 

Wing membrane black except for reddish orange skin along digits; dorsal pelage reddish orange; 

forearm 52-53 mm; skull as in Figure 55 Myotis rufopictus 

Wings uniformly colored; dorsal pelage gray or brown 10 

Posterior margin of wing membrane terminates on ankle (fig. 16A); hind foot 15-17 mm; forearm 

44-49 mm; skull as in Figure 53 Myotis macrotarsus 

Posterior margin of wing membrane terminates on side of foot, below ankle (figs. 16B,C); hind foot 

6-12 mm; forearm 30-38 mm 11 

Posterior margin of wing membrane terminates on side of foot, at least 1 mm above base of 
outermost toe (fig. 16B); hind foot 10-12 mm; forearm 35-38 mm; skull as in Figure 52 

Myotis horsfieldii 

Posterior margin of wing membrane terminates on side of foot at base of outermost toe (fig. 1 6C); 

hind foot 6-8 mm; forearm 30-34 mm; skull as in Figure 54 Myotis muricola 

Pads present on feet and wrists (fig. 1 7); forearm 22-30 mm 13 

No pads on feet and wrists; forearm 30-55 mm 15 

Pads on feet and wrists pink or white; dorsal fur dark at base; posterior margin of wing membrane 



20 FIELDIANA: ZOOLOGY 



B 





Fig. 14. Ventral views of muzzles of Murina (A) and Myotis (B). 



terminates on side of foot, at base of outermost toe; five pairs of upper cheek teeth; forearm 26- 

30 mm Glischropus tylopus 

1 3'. Pads on feet and wrists dark brown; dorsal fur not dark at base; posterior margin of wing membrane 
terminates on ankle or on side of foot, at least 1 mm above base of outermost toe; four pairs of 
upper cheek teeth; forearm 22-27 mm (Tylonycteris) 14 

14. Lower third of posterior margin of ear distinctly thicker than rest of ear (fig. 18 A); forearm 24-27 
mm Tylonycteris robustula 

14'. Lower third of posterior margin of ear not distinctly thicker than rest of ear (fig. 18B); forearm 22- 
25 mm Tylonycteris pachypus 

15. Most distal bone of third digit of wing about three times length of adjoining phalanx 

(Miniopterus) 1 6 

15'. Most distal bone of third digit of wing (fig. 2) not more than two times length of adjoining phalanx 
17 

16. Forearm 5 1-55 mm; tail 50-61 mm; skull as in Figure 50 Miniopterus tristis 

16'. Forearm 42-46 mm; tail 46-57 mm; skull as in Figure 49 Miniopterus schreibersi 

16*. Forearm 34-39 mm; tail 34-43 mm; skull as in Figure 48 Miniopterus australis 

17. Posterior margin of wing membrane terminates on lower leg, at or above ankle; fifth digit of wing 
(see fig. 2) does not extend beyond midpoint between first and second joint of third digit .... 18 

17'. Posterior margin of wing membrane terminates on side of foot, below ankle (as in figs. 16B,C); fifth 
digit of wing extends beyond midpoint between first and second joint of third digit 19 

1 8. Inner upper incisors transversely elongate in cross-section, about three times as wide as outer upper 
incisors; outer secondary cusp on inner upper incisors is not as high but almost as wide as the main 
cusp; forearm 34-39 mm; skull as in Figure 56 Philetor brachypterus 

18'. Inner upper incisors conical in shape, about two times as wide as outer upper incisors; outer 





Fig. 15. Left external ears (pinnae) of some vespertilionids. A, Murina/ Harpiocephalus; B, Kerivoula/Phoniscus; 
C, Myotis; D, Pipistrellus. 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



21 




Fig. 1 6. Left hind feet of Myotis spp., showing attachment of wing membrane. A, hi. macrotarsus; B, M. horsfieldii; 
C, M. muricola. 

secondary cusp on inner upper incisors is much narrower than main cusp; forearm 38^40 mm . . 

Pipistrellus stenopterus 

1 9. Forearm about 39 mm Pipistrellus petersi 

19'. Forearm 32-36 mm; tail 29-34 mm; skull as in Figure 57 Pipistrellus javanicus 

19". Forearm 30-32 mm; tail 26-31 mm; skull as in Figure 58 Pipistrellus tenuis 



Family Molossidae: 
Free-Tailed Bats 

This family of at least 80 species is represented 
in the Philippines by only four species. As the 
English name for the family implies, molossid bats 
are the only bats with long tails that project well 
beyond the posterior edge of the interfemoral 
membrane (fig. 7E). The wings tend to be unusu- 
ally long and narrow for the size of the bats. There 
is no noseleaf; the lips are often wrinkled and thick 
and the upper lip thickly sprinkled with short, bris- 
tly hairs (fig. 8E). The eyes are moderate to small. 
The external ear is moderately small, with the tra- 
gus small and the antitragus well developed. Phil- 
ippine molossids are of moderate to large size, 
with forearms ranging from 40 to 90 mm (table 



5). Two of the species in the Philippines (both 
Cheiromeles) have only scattered hairs over most 
of their bodies, leading to their common name, 
"naked bats." 

The skulls of molossids are characterized by 
moderately sturdy construction, lack of postor- 
bital processes, and broad, strong molars (fig. 60). 
Among Philippine molossids, the anterior edge of 
the palate is continuous, so that the upper pairs 
of incisors are separated by no more than a narrow 
gap. The jaws are powerful, and the large masse- 
teric muscles on the head attach to a moderate to 
high sagittal crest. 

These bats are strong, fast fliers that feed pri- 
marily on beetles and other large insects. Chaere- 
phon plicata, which roosts in caves and occasion- 






Fig. 1 7. Pad on right wrist of Glischropus. 



Fig. 1 8. Left ears of Tylonycteris. A, T. robustula; B, 
T. pachypus. 



22 



FIELDIANA: ZOOLOGY 



ally in buildings, probably once formed the largest 
colonies of bats in the Philippines, exceeding 
100,000 individuals, according to early descrip- 
tions, but most colonies are now much reduced in 
size or entirely destroyed. Cheiromeles parvidens 
and C. torquatus are known to roost in hollow 



coconut palms, hollow trees, and occasionally in 
caves, and Mops sarasinorum in hollow trees, where 
they form colonies of four to hundreds of indi- 
viduals. 

The systematics of bats of this family has been 
reviewed by Freeman (1981). 



Key to Molossidae 

1. Body furred; upper lip wrinkled; forearm 39^45 mm; ears joined together over top of head by 
narrow band of skin 2 

1'. Body almost naked; upper lip not wrinkled; forearm 73-90 mm; ears separate, not joined across 
top of head (Cheiromeles) 3 

2. Five upper cheek teeth, including a small anterior premolar that is very much smaller than the 
other cheek teeth (fig. 60); condylocanine length 14.9-15.5 mm; forearm 40-43 mm; skull as in 
Figure 60 Chaerephon plicata 

2'. Four upper cheek teeth; condylocanine length 17.3-17.8 mm; forearm 39-45 mm 

Mops sarasinorum 

3. Forearm 73-78 mm (occurs throughout the Philippines except Palawan) Cheiromeles parvidens 

3'. Forearm 80-90 mm (occurs only in Palawan region) Cheiromeles torquatus 



Notes 

The following brief comments on the status of 
several species of Philippine bats are intended to 
make users of the key aware of potential taxonom- 
ic problems and of recent records of bats that are 
referred to in the text. 



Acerodon lucifer 

This species is known only from the type series 
collected on Panay Island in 1888. Based on cur- 
rent knowledge, it differs from most Acerodon ju- 
batus only on the basis of smaller size, but it over- 
laps with some populations for all measurements. 
It is possible that careful study will show it to have 
denning characters that are not yet recognized, or, 
alternatively, to be a geographic variant of the 
widespread A. jubatus. 



Cynopterus brachyotis 

Philippine representatives of the genus Cynop- 
terus have until recently been referred to C. 
brachyotis, which is widespread in Southeast Asia. 
Based primarily on morphometric analyses, how- 
ever, Kitchener and Maharadatunkamsi (1990) 
split what had been known as Cynopterus brachy- 



otis into several species, allocating Philippine 
specimens to C luzoniensis based on a single sam- 
ple from Negros Island. We have not yet been able 
to examine enough material to evaluate their pro- 
posed changes and have therefore followed the 
older terminology, while recognizing that further 
study may support the proposed revision. 



Pteropus dasymallus 

This species has been reported previously only 
from Japan, Taiwan, and adjacent and intervening 
islands. We have recently identified several spec- 
imens in fmnh, pnm, and usnm from the Babuyan 
and Batanes islands, which lie between northern 
Luzon and Taiwan, as representing this species. 
Details will be published elsewhere. The species 
is very similar to P. hypomelanus, differing most 
visibly in that the pelage is longer and denser, and 
the dorsal surface of the hind leg is thickly furred 
nearly to the ankle (rather than being nearly na- 
ked). 



Hipposideros coronatus 

No specimens of this moderately large horse- 
shoe bat have been reported since the species was 
described in 1871 (Peters, 1871). Our placement 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



23 



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of the species in the key is based solely on the 
original description and the comments of Taylor 
(1934). 



Rhinolophus anderseni 

This species is known only from the holotype, 
which we have been unable to examine. As noted 
under the following species, it is possible that re- 
cently taken specimens may be referred to this 
species, but this remains uncertain. 

Rhinolophus arcuatus 

Heaney et al. (1991) recently noted that bats 
previously identified as belonging to this species 
appear to fall into two morphotypes: one is slightly 
smaller with a narrower anterior noseleaf and is 
typically found in lowland caves or disturbed hab- 
itats, and the other is slightly larger with a pro- 
portionately wider anterior noseleaf and is usually 
found in primary upland forest. We designate these 
in the key as "i?. arcuatus-s" and "R. arcuatus- 
1", respectively. J. Hill (in litt.) has suggested that 
one of these may be R. anderseni, but we have not 
examined the relevant holotypes and so remain 
uncertain of the appropriate use of names. More 
specimens, especially of the larger upland form, 
are badly needed. 

Rhinolophus inops and 
R. subrufus 

Our studies of specimens currently in collections 
have indicated that Philippine bats in this species 
group may be broken into at least two morphs, 
based on size; these are very similar in morphology 
of the noseleaf and skull. In this key, we refer the 
range of smaller bats to R. inops and that of the 
larger ones to R. subrufus. However, we note that 
there are several unresolved problems with these 
bats. 

First, the noseleaf of the holotype of R. inops 
has a distinctly shaped sella: the tip is modified 
into a downward-projecting triangular pouch (An- 
dersen, 1905c). This modified sella is not found 
in any other specimen that has been referred to R. 
inops or in any other species in the genus; it may 
represent an aberration found in only one indi- 
vidual or it may be a diagnostic character for a 
species still known only from the holotype, with 
the other specimens referred to R. inops actually 
representing an undescribed species. 



24 



FIELDIANA: ZOOLOGY 



Second, the definition of/?, inops is complicated 
by the variation in size among specimens that we 
have assigned to R. inops; available specimens 
from Negros are consistently smaller than those 
from Leyte and Biliran, whereas those from Ca- 
tanduanes are intermediate (specimens from else- 
where are inadequate for comparison). It is pos- 
sible that each morph represents a distinct species. 
Additional specimens and further study will be 
needed to clarify the status of all of these bats. 



Rhinolophus macron's 

The Philippine form of R. macrotis was first 
described as a distinct species, R. hirsutus (An- 
dersen, 1905b), but was later subsumed under R. 
macrotis (Tate, 1 943). We have examined the ho- 
lotypes and referred specimens of both taxa and 
find that they differ in overall size, in proportion- 
ate tail length, and in the size and shape of noseleaf 
structures, particularly the sella. We suspect that 
the Philippine population is morphologically dis- 
tinct and genetically independent and will even- 
tually be shown to be a distinct species, but we 
refrain from making this change because we have 
not conducted comprehensive studies. 



Harpiocephalus harpia 

Three specimens of this large and striking ves- 
pertilionid have been taken recently, one on Leyte 
(Rickart et al., in prep.), one on Luzon (Heaney et 
al., in prep.), and one on Negros (Utzurrum, pers. 
comm.). Further details will be published else- 
where. 



Acknowledgments 

We wish to thank the curators and staff of the 
American Museum of Natural History, British 
Museum (Natural History), Delaware Museum of 
Natural History, Philippine National Museum, 
Royal Ontario Museum, Silliman University Mu- 
seum of Natural History, University of Michigan 
Museum of Zoology, and United States National 
Museum of Natural History for allowing us to 
examine specimens in their care. We are partic- 
ularly grateful to G. B. Corbet, P. C. Gonzales, P. 
Jenkins, K. F. Koopman, and W. T. Stanley for 
assistance with access to specimens. We have re- 



ceived an exceptional amount of enthusiastic as- 
sistance with collecting bats in the field; although 
we are unable to mention all of those who have 
generously given us help, we must acknowledge 
the efforts of D. Balete, R. Fernandez, P. C. Gon- 
zales, P. D. Heideman, A. Manamtam, E. A. Rick- 
art, and R. C. B. Utzurrum. We are grateful to the 
Protected Areas and Wildlife Bureau of the Phil- 
ippines, especially J. Caleda, C. Custodio, W. Dee, 
M. Mendoza, and S. Penafiel for their continuing 
support and cooperation. The illustrations of skulls 
and the map were prepared by T. B. Griswold; all 
others were prepared by J. Sedlock, whom we es- 
pecially thank for her efforts as a volunteer. C. M. 
Francis helped NRI distinguish between Philip- 
pine bats that also occur in Borneo. D. Balete, S. 
M. Goodman, K. F. Koopman, H. Miranda, A. 
T. Peterson, E. A. Rickart, W. Schutt, D. Willard, 
and an anonymous reviewer kindly reviewed the 
manuscript and provided helpful comments. We 
especially thank K. F. Koopman for his highly 
constructive input into every stage of this study. 
This research has been generously supported by 
the U.S. National Science Foundation (BSR- 
8514223) and the MacArthur Foundation (90- 
9272). The Dee Fund of the fmnh provided travel 
funds to NRI. An earlier draft of this manuscript 
formed part of a master's thesis by NRI at Cornell 
University. 



Literature Cited 

Andersen, K. 1905a. On some bats of the genus Rhi- 
nolophus, with remarks on their mutual affinities, and 
descriptions of twenty-six new forms. Proceedings of 
the Zoological Society of London, 2: 75-145. 

. 1905b. On the hats of the Rhinolophus philip- 

pinensis group, with descriptions of five new species. 
Annals and Magazine of Natural History, 7th series, 
16: 243-256. 

-. 1905c. On the bats of the Rhinolophus arcuatus 



group, with descriptions of five new forms. Annals and 
Magazine of Natural History, 7th series, 16: 281-288. 
1 905d. On the bats of the Rhinolophus macro- 



tis group, with descriptions of two new forms. Annals 
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1912. Catalog of the Chiroptera in the collec- 



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Anthony, E. L. P. 1988. Age determination in bats, 
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Harrison, J. 1966. An Introduction to the Mammals 
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Hauge, P., J. Terborgh, B. Winter, and J. Parkinson. 

1986. Conservation priorities in the Philippine Ar- 
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. 1991. An analysis of patterns of distribution 

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Heaney, L. R., P. C. Gonzales, and A. C. Alcala. 

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Heaney, L. R., and P. D. Heideman. 1987. Philippine 
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Heaney, L. R., and R. L. Peterson. 1984. A new 
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26 



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INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



27 




1 



as 



28 



FIELDIANA: ZOOLOGY 





Fig. 20. Skull of Cynopterus brachyotis (ummz 156664). Scale = 10 mm. 




Fig. 21. Skull of Dobsonia chapmani (dmnh 5131). Scale = 10 mm. 





Fig. 22. Skull of Eonycteris robusta (ummz 162221). Scale = 10 mm. 
INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



29 





Fig. 23. Skull of Eonycteris spelaea (ummz 158482). Scale = 10 mm. 






Fig. 24. Skull of Haplonycteris fischeri (ummz 157047). Scale = 10 mm. 






Fig. 25. Skull of Harpyionycteris whiteheadi (ummz 158838). Scale = 10 mm. 



30 



FIELDIANA: ZOOLOGY 






Fig. 26. Skull of Macroglossus minimus (ummz 158725). Scale = 10 mm. 





Fig. 27. Skull of Nyctimene rabori (usnm 458281). Scale = 10 mm. 






Fig. 28. Skull of Ptenochirus jagori (ummz 158513). Scale = 10 mm. 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



31 




Fig. 29. Skull of Pteropus hypomelanus (usnm 105443). Scale = 10 mm. 






Fig. 30. Skull of Pteropus pumilus (ummz 158517). Scale = 10 mm. 



32 



FIELDIANA: ZOOLOGY 




Fig. 31. Skull of Pteropus vampyrus (ummz 158849). Scale = 10 mm. 




Fig. 32. Skull of Rousettus amplexicaudatus (ummz 162304). Scale = 10 mm. 






Fig. 33. Skull of Emballonura alecto (usnm 458536). Scale = 10 mm. 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



33 







Fig. 34. Skull of Saccolaimus saccolaimus (usnm 458555). Scale = 10 mm. 




Fig. 35. Skull of Taphozous melanopogon (usnm 458571). Scale = 10 mm. 



34 



FIELDIANA: ZOOLOGY 




Fig. 36. Skull of Megaderma spasma (ummz 160295). Scale = 10 mm. 






Ate&i^ 



Fig. 37. Skull of Hipposideros ater (rom 40735). Scale = 10 mm. 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



35 








Fig. 38. Skull of Hippos ideros diadema (ummz 160296). Scale = 10 mm. 






Fig. 39. Skull of Hipposideros obscurus (su 787). Scale = 10 mm. 



36 



FIELDIANA: ZOOLOGY 






^feb^^l 



Fig. 40. Skull of Hipposideros pygmaeus (ummz 156872). Scale = 10 mm. 




Fig. 41. Skull of Rhinolophus arcuatus-X (ummz 158526). Scale = 10 mm. 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



37 






Fig. 42. Skull of Rhinolophus arcuatus-s (ummz 157106). Scale = 10 mm. 




Fig. 43. Skull of Rhinolophus inops (usnm 459495). Scale = 10 mm. 



38 



FIELDIANA: ZOOLOGY 





Fig. 44. Skull of Rhinolophus macrotis (ummz 160328). Scale = 10 mm. 




Fig. 45. Skull of Rhinolophus philippinensis (ummz 459497). Scale = 10 mm. 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



39 









Fig. 46. Skull of Rhinolophus virgo (usnm 459454). Scale = 10 mm. 






Fig. 47. Skull of Kerivoula hardwickii (usnm 45951 1). Scale = 10 mm. 






Fig. 48. Skull of Miniopterus australis (usnm 458658). Scale = 10 mm. 



40 



FIELDIANA: ZOOLOGY 






Fig. 49. Skull of Miniopterus schreibersi (ummz 157001). Scale = 10 mm. 




Fig. 50. Skull of Miniopterus tristis (usnm 458680). Scale = 10 mm. 




Fig. 51. Skull of Murina cyclotis (usnm 573776). Scale = 10 mm. 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



41 






Fig. 52. Skull of Myot is horsfieldii (ummz 158872). Scale = 10 mm. 





Fig. 53. Skull of Myot is macrotarsus (ummz 162364). Scale = 10 mm. 






Fig. 54. Skull of Myotis muricola (ummz 158873). Scale = 10 mm. 



42 



FIELDIANA: ZOOLOGY 




Fig. 55. Skull of Myot is rufopictus (ummz 158878). Scale = 10 mm. 







Fig. 56. Skull of Philetor brachypterns (usnm 573781). Scale = 10 mm. 






Fig. 57. Skull of Pipistrellus javanicus (usnm 459762). Scale = 10 mm. 



INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 



43 






Fig. 58. Skull of Pipistrellus tenuis (usnm 160338). Scale = 10 mm. 




Fig. 59. Skull of Scotophilus kuhlii (usnm 458704). Scale = 10 mm. 




Fig. 60. Skull of Chaerephon plicata (usnm 304238). Scale = 10 mm. 



44 



FIELDIANA: ZOOLOGY 



HECKMAN 

BINDERY INC. 

JAN 95 

^.T„.^N N MANCHESTER.